Anais da Academia Brasileira de Ciências (2018) 90(3): 2865-2872 (Annals of the Brazilian Academy of Sciences) Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 http://dx.doi.org/10.1590/0001-3765201820170715 www.scielo.br/aabc | www.fb.com/aabcjournal

Acanthagrion marinae sp. nov. (Zygoptera: ): a new species of the apicale group

FEDERICO LOZANO1 and MARCIEL E. RODRIGUES2

1Laboratorio de Biodiversidad y Genética Ambiental/BioGeA, Universidad Nacional de Avellaneda, Mario Bravo 1460 esq. Isleta, 1870 Piñeyro, Buenos Aires, Argentina 2Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, Km 16, 45662-900 Ilhéus, BA, Brazil

Manuscript received on September 11, 2017; accepted for publication on February 5, 2018

ABSTRACT Acanthagrion marinae sp. nov. (Holotype male: BRASIL, Mato Grosso do Sul, Campo Grande, UFMS, 20°29’56.26’’S - 54°36’48.43’’W, 547m, leg. M.E. Rodrigues, 03.ii.2015, collection code: VZYG437, MLP) is described and illustrated on the basis of 15 males. The new species belongs to the apicale species group by having horns on S10 and sclerotized hooks on tip of distal segment of the genital ligula. It can be easily distinguished from other species of the group by a combination of characters of the genital ligulae (presence of setae on segment two; absence of setae at flexure; distal lateral lobes of segment three absent). Notes on habitat and a modification of previous keys for the species of theapicale group are provided. Key words: Acanthagrion, Coenagrionidae, Neotropics, , Zygoptera.

INTRODUCTION cerci decumbent forming an angle of 45° with posterior margin of S10, with dorso-basal tubercle; The neotropical Acanthagrion was erected females without a well defined mesepisternal in 1876 by Baron Edmond Selys-Longchamps to carina between mesostigmal plates and medio- include nine species of Coenagrionidae. Currently dorsal carina, generally with mesepisternal fossae it includes 44 coenagrionids ranging from Texas on the dorsum of the pterothorax and vulvar spine (USA) to Buenos Aires (Argentina) (Garrison on abdominal segment S8. et al. 2010, Machado 2012). According to von Leonard (1977) provided the first revision of Ellenrieder and Lozano (2008) and Garrison et the genus in which he defined nine species groups. al. (2010) species of Acanthagrion are small to The three species originally included within the medium size with wings generally hyaline, CuP apicale species group (A. apicale, A. obsoletum, of HW reaching posterior margin of wing; flexure and A. phallicorne) share the presence of horns of male genital ligula short with lateral lobes of in S10 of males, elevated mesepisternal fossae in segment 3, when present, distal to flexure; male females, and presence of orange coloration in head

Correspondence to: Federico Lozano and thorax of mature specimens. Acanthagrion E-mail: [email protected] apicale and A. obsoletum also share the presence

An Acad Bras Cienc (2018) 90 (3) 2866 FEDERICO LOZANO and MARCIEL E. RODRIGUES of sclerotized hooks on either side of the tip Westfall and May (2006). Color pattern of head and of distal segment of genital ligula. Muzón and abdomen follows Lozano (2013). Lozano (2005) described A. hartei from Ecuador Collection acronyms used in the text: and included it within the apicale group, based BioGeA: Laboratorio de Biodiversidad y on shared characters with its members. Recently Genética Ambiental, Avellaneda, Argentina Machado (2012) provided a revision of the group MLP: Departamento Científico Entomología, and added four new species (A. chicomendesi, A. Museo de La Plata, La Plata, Argentina flaviae, A. kaori, and A. triangulare), and accorded UFMS: Universidade Federal de Mato Grosso specific rank for Fraser’s A. apicale descendens, do Sul, Mato Grosso do Sul, Brazil bringing the total number of species of the group UFSCAR: Universidade Federal de São to nine. Carlos, São Paulo, Brazil In this contribution a new species of the Measurements are given in mm. A list of apicale species group from Brazil and Paraguay is examined material is provided; country, department, described, diagnosed and illustrated. state, province or any other political/administrative The ZooBank Life Science Identifer (LSID) equivalent subdivision, locality (names of localities of this publication is: urn:lsid:zoobank.org:pub: were transcribed from the label and are not translated 0951B791-04E1-41E5-8731-E71E3D57CD96. to English), geographic coordinates and altitude, collector/s, date of collection, number and sex of MATERIALS AND METHODS specimens, and deposition of material are indicated. When geographic coordinates were not included Specimens were studied with the aid of a Leica in the original label, they were culled from the MS5 stereomicroscope in the Laboratorio de Global Gazetteer website (http://www.fallingrain. Biodiversidad y Genética Ambiental (BioGeA) com/world/) with the aid of Google Earth version of the Universidad Nacional de Avellaneda 7.1.7.2606, and these data are enclosed in braces ({ }). (UNDAV). Illustrations were made with the aid of The map was created electronically using QGIS digital camera coupled to the stereomicroscope and version 2.16.3. an open-source design program (Inkscape version 0.91. at www.inkscape) and are not to scale. RESULTS Specimens for SEM were cleaned in acetone, air Acanthagrion marinae sp. nov. dried and mounted on SEM stubs with carbon- ZooBank Life Science Identifer (LSID) - conductive adhesive tabs, and then sputter coated urn:lsid:zoobank.org:act:D1F0D354-41FC-444A- with gold/ palladium alloy and examined with a B57B-71CBA362904C Jeol JSM 6360 LV scanning electron microscope Types. Holotype ♂: BRASIL, Mato Grosso do in the Museo de La Plata, Buenos Aires, Argentina. Sul, Campo Grande, UFMS, 20°29’56.26’’S - Wing venation follows Riek and Kukalová- 54°36’48.43’’W, 547m, leg. M.E. Rodrigues, Peck (1984), modified by Bechly (1996). 03.ii.2015, MLP. Paratypes 14 ♂: BRASIL, Abbreviations for structures used throughout the Mato Grosso do Sul: Campo Grande, Anhanduí, text are as follows: FW: forewing; HW: hindwing; 20°30’24.18’’S - 54°38’28.60’’W, 507m, leg. pt: pterostigma; Ax: antenodals; Px: postnodals; M.E. Rodrigues, 24.xi.2014, 3♂, UFSCAR; pt: pterostigma. S1–10: abdominal segments 1 to Campo Grande, Brejo bonito, 20°32’05.96’’S 10. Color pattern terminology of thorax follows - 54°44’52.91’’W, 493m, leg. M.E. Rodrigues,

An Acad Bras Cienc (2018) 90 (3) Acanthagrion marinae sp. nov. 2867

15.xi.2014, 1♂, MLP; Campo Grande, Córrego segment 2 is only shared with A. phallicorne; Bandeira, 20°29’33.68’’S - 54°34’44.50’’W, 598m, from which it can be easily separated because A. leg. L.C. Alves, 16.i.2015, 1♂, MLP; Campo marinae lacks setae on flexure. Tip of segment 3 Grande, Cotia, 20°32’37.78’’S - 54°41’29.08’’W, with sclerotized hooks on either side. 508m, leg. L.C. Alves, 22.i.2015, 3♂, MLP; Description of holotype male (Fig. 1): Campo Grande, Lageado, 20°32’26.99’’S - Head. Labrum light blue postero-median spot 54°38’23.64’’W, 498m, leg. L.C. Alves, 20.i.2015, black; anteclypeus light blue; postclypeus light blue 1♂, MLP; Campo Grande, Morada do sol, with two black spots and posterior margin black; 20°39’18.94’’S - 54°21’59.53’’W, 544m, leg. antefrons light blue; epicranium mostly black E.B. Moura, 16.i.2015, 1♂, MLP; Campo Grande, with two greenish light blue spots behind scape, Norberto, 20°34’16.61’’S - 54°36’31.33’’W, a very diffuse greenish spot next to each ocellus, 536m, leg. M.E. Rodrigues, 24.xi.2014, 2♂, a T-shaped spot within ocellar triangle; occipital BioGeA; Campo Grande, Prainha, 20°33’57.66’’S bar greenish light blue; antennifer and scape - 54°30’21.21’’W, 602m, leg. M.E. Rodrigues, anteriorly light blue, posteriorly black; reminder 02.ii.2015, 1♂, MLP. PARAGUAY, Cerro Corá, of the antenna missing; postocular spots greenish {22°37’00’’S - 55°59’00’’W, 266m}, leg. L.A. light blue, subrectangular, approximately equal Bulla, 30.xi.1973, 1♂, MLP. in size to ocellar triangle; most posterior point of Specimens examined. 15♂ head located at compound eyes; occipital area pale brown, except for two dark brown rounded spots on ETYMOLOGY each side of occipital foramen. The name marinae (Latinized genitive feminine Prothorax. Anterior lobe with central spot noun) pays homage to the memory of Marina dos greenish light blue occupying almost all of dorsum, Santos Rodrigues, mother of Marciel E. Rodrigues remainder of dorsum black, mostly black laterally one of the authors of this article. except for narrow anterior greenish light blue spot; middle lobe entirely black, dorsolateral spots DIAGNOSIS greenish light blue, propleuron greenish light blue; Pale color areas of head and thorax are light blue; posterior lobe black with greenish light blue lateral this can easily differentiate A. marinae from most streaks; lateral margins rounded lacking a medial other species included within the apicale group projection. (except for A. descendens and some specimens of Pterothorax (Fig. 1a). Predominantly greenish A. obsoletum). However, coloration has proven light blue. Mid-dorsal black stripe not interrupted at not to be a reliable character since other orange middorsal carina; antealar sinus black; antehumeral species within Acanthagrion (A. obsoletum and A. ascendens) have both morphs (Machado 2012). Acanthagrion marinae has horns on S10 with tips rounded, directed dorso-posteriorly and not surpassing distal margin of S10; this combination of characters is unique within the apicale group. Cerci with inner margin straight longer than the length of S10 can easily differentiate this species Figure 1 - a. Holotype, general view; b. Holotype, terminalia from A. obsoletum. Genital ligulae with setae on lateral view.

An Acad Bras Cienc (2018) 90 (3) 2868 FEDERICO LOZANO and MARCIEL E. RODRIGUES stripe light greenish blue, uninterrupted but not between Px3 and Px4; IR1 beginning at Px8; five reaching antealar crest; anterior to posterior limit of cells posterior to pt. mesinfraepisternum antehumeral stripe widening Abdominal segments. S1: tergum: posterior slightly; humeral stripe black; interpleural margin of anterior spot in contact with posterior suture black; incomplete stripe on metapleural stripe of S1; posterior stripe complete, its central suture pale brown; mesinfraepisternum black region in contact with posterior margin of tergum; with postero-ventral angle greenish light blue; lateral margins in contact with ventrolateral margins of tergum in lateral view; sternum: pale brown, metinfraepisternum pale greenish light blue; with a triangular black spot on anterior margin; venter of thorax pale brown without black spots; S2: tergum: dorsal spot subrectangular, widening mesostigmal plates without diagonal carina, with posteriorly; anterior margin in contact with anterior inner half black and outer half light blue, without margin of tergum; posterior margin in contact with modifications on posterior angle; interlaminar sinus posterior stripe of S2; posterior stripe in contact black; anterior margin of sinus slightly convex; with posterior margin of tergum; lateral margins lateral tips rounded not projected anteriorly. reaching halfway down terga in lateral view; Legs. Coxae and trochanters pale brown; sternum: anterior lamina and posterior hamuli pale femora with extensor margin black, flexor margin brown; anterior hamuli greenish light blue; S3 to pale brown; tibiae with extensor margin pale S6: terga: dorsal T-shaped spots reaching anterior brown except for a longitudinal row of elongated margin through thin black line; anterior margins black spots just behind row of external spurs, blunt; lateral arms directed anteriorly in S3 and flexor margin black, femoral spurs shorter than S4, reaching halfway down terga in lateral view; intervening spaces, tibial spurs equal to intervening posterior stripes visible in S3 and S4; sterna: pale spaces; tarsi pale brown with tips black. brown with black midventral line; S7: tergum: Wings (Fig. 2). CuP reaching posterior margin subapical row of spines present; with a black of wing; arculus opposite Ax2. FW left: Px 11; RP2 subrectangular spot that occupies the anterior two beginning between Px4 and Px5; IR1 beginning thirds of tergum; posterior margin with a triangular at Px8; four cells posterior to pt. FW right: Px 10; concavity posterior third light blue, except for a RP2 beginning at Px5; IR1 beginning at Px8; five thin black spot posterior to subapical row of spines; sternum: pale brown with black midventral stripe; cells posterior to pt. HW left: Px 9; RP2 beginning S8 and 9 (Fig. 1b): Light blue except for a thin black between Px3 and Px4; IR1 beginning at Px7; five spot posterior to subapical row of spines; subapical cells posterior to pt. HW right: Px 9; RP2 beginning row of spines present; sternum: pale brown with black midventral stripe; S10 (Figs. 1b, 3c): tergum trapezoidal in lateral view; posterior margin with horns that do not surpass posterior margin of S10 in lateral view; tergum black with a mid-dorsal reniform light blue spot; venter pale brown. Height of S10 / height of S9 1.32. Genital ligula (Fig. 3a-b). Segment 2 with setae on lateral margins; flexure shorter than segment 3, without setae. Segment 3 with lateral Figure 2 - Holotype, wings. lobes located at 0.5 the length of segment 3;

An Acad Bras Cienc (2018) 90 (3) Acanthagrion marinae sp. nov. 2869

Figure 3 - a. Paratype Brazil, genital ligula ventral view; b. Paratype Brazil, genital ligula lateral view; c. Paratype Brazil, male terminalia lateral view; d. Paratype Brazil, male terminalia posterior view. ental surface transversal fold between lateral MEASUREMENTS lobes and apicolateral lobes (TL and ML sensu Head: max. length 0.94; width between compound Machado 2012); distal margin without indentation; eyes along anterior margin 1.37. Legs: femur 1 apicolateral lobes with sclerotized hook. length 1.37; femur 2 length 1.75; femur 3 length Cerci (Fig. 3c-d). Black with inner surface 2.22. Wings: FW length left 16.81, right 16.94; slightly paler. Basal tubercle rounded. Inner HW length left 15.69, right 15.69. Abdomen: margin in dorso-medial view almost straight; tips max. length 24.91; S1 max. length 0.57; S2 max. rounded. Dorsal margin in lateral view with two length 1.37; S3 max. length 4.11; S4 max. length 4.49; S5 max. length 4.58; S6 max. length 4.23; constrictions. S7 max. length 3.31; S8 max. length 1.37; S9 max. Paraprocts (Fig. 3c-d). Base pale brown; length 0.76; S10 max. length 0.47; S9 height 0.90; apophysis black with tips acute, in lateral view with S10 height 1.18. Cerci: length: 0.94; distance tips directed upwards, in ventral view tips directed surpassing posterior margin of S10 in lateral view inwards. Paraprocts not surpassing tip of cerci. 0.38. Paraprocts: distance surpassing posterior

An Acad Bras Cienc (2018) 90 (3) 2870 FEDERICO LOZANO and MARCIEL E. RODRIGUES margin of S10 in lateral view 0.33. Total length max. length 4.40 [3.92–4.91]; S6 max. length 4.15 30.26. [3.78–4.63]; S7 max. length 3.14 [2.83–3.40]; S8 max. length 1.34 [1.18–1.42]; S9 max. length 0.74 VARIATIONS IN PARATYPES [0.70–0.80]; S10 max. length 0.48 [0.40–0.57]; S9 Little variation was observed in the paratypes; height 0.98 [0.87–1.09]; S10 height 1.21 [1.10– minor differences in the extent of black color of 1.32]. Cerci: length: 0.94 [0.90–1.04]; distance head and thorax probably are due to age differences. surpassing posterior margin of S10 in lateral view No differences were observed in genital ligula or 0.40 [0.33–0.52]. Paraprocts: distance surpassing male terminalia. Wings: CuP reaching posterior posterior margin of S10 in lateral view 0.39 [0.29– margin of wing or CuP&AA’; arculus opposite Ax2 0.47]. Total length 29.42± 1.56 [26.90–31.63]. or slightly distal to it. FW left: Px 10 (60%) or 11 FEMALE (40%); RP2 beginning between Px4 and Px5 (50%) or at Px5 (50%); IR1 beginning at Px7 (30%) or at Despite surveys in the area where males were Px8 (70%); four (70%) or five (30%) cells posterior collected no female has been found. Further to pt. FW right: Px 10 (50%) or 11 (50%); RP2 research is needed in order to collect and describe beginning between Px4 and Px5 (50%) or at Px5 the female.

(50%); IR1 beginning at Px7 (30%), at Px8 (60%) HABITAT or at Px9 (10); three (10%), four (40%), five (30%) or six (20%) cells posterior to pt. HW left: Px 8 Acanthagrion marinae has been recorded for Brazil (30%) or 9 (70%); RP2 beginning between Px3 and and Paraguay (Fig. 4a). The specimens from Brazil Px4 (40%) or at Px4 (60%); IR1 beginning at Px7 were collected in the Veredas or palm swamps (100%); five (80%) or 6 (20%) cells posterior to areas which are hygrophilous communities with pt. HW right: Px8 (30%), 9 (60%) or 10 (10%); an enormous floristic and structural complexity RP2 beginning between Px3 and Px4 (60%), at dominated by Mauritia flexuosa L. (Arecaceae). Px4 (30%) or between Px4 and Px5 (10%); IR1 The Veredas feature special importance for water beginning at Px6 (10%), between Px6 and Px7 and biodiversity maintenance in the Cerrado (10%) or at Px7 (80%); four (40%), five (50%) or biome (Alencar Fagundes and Ferreira 2016). six (10%) cells posterior to pt. The Brazilian Forest Code considers the Veredas as a permanent preservation zone (Law number MEASUREMENTS (N=10) 12.651/2012) due to the ecological services they Head: max. length 0.89 [0.80–0.94]; width provide. The Brazilian specimens were collected in between compound eyes along anterior margin urban and rural places of the municipality of Campo 1.34 [1.23–1.42]. Legs: femur 1 length 1.30 [1.18– Grande, MS. The males were found perching on the 1.42]; femur 2 length 1.75 [1.56–1.84]; femur 3 vegetation along the stream banks or on emergent length 2.12 [1.95–2.36]. Wings: FW length left twigs within the streams (Fig. 4b), usually in sunlit 16.43 [15.44–18.06], right 16.42 [15.69–17.81]; clearings with dense canopy. No information is HW length left 15.42 [14.57–17.06], right 15.62 available for specimens collected in Paraguay. [14.82–16.94]. Abdomen: max. length 24.31 DISCUSSION [21.67–26.40]; S1 max. length 0.64 [0.60–0.71]; S2 max. length 1.33 [1.23–1.42]; S3 max. length 4.07 Acanthagrion marinae belongs to the apicale [3.73–4.44]; S4 max. length 4.39 [4.02–4.82]; S5 species group, since it has horns on S10 and there

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Figure 4 - a. Distribution map; b. Male perching on twig. Photo: Thiago Henrique Stefanello. are sclerotized hooks on tip of distal segment. No 3’ Penis segment 2 with no patch of setae ... 4 orange coloration has been observed so far in the (See Machado 2012) specimens of A. marinae. The presence of setae on 9. Flexure of genital ligula with setae ... A. segment two is only shared with A. phallicorne, phallicorne 9’ Flexure of genital ligula without setae ... A. from which can be easily differentiated because A. n. sp phallicorne has setae on the flexure of the genital ligulae and there are distal lateral lobes on the ACKNOWLEDGMENTS segment three of the penis. We thank Rosser Garrison and Javier Muzón for the According to the key provided by Machado critical reading of the manuscript; and Emanuelle (2012), males of A. marinae run to couplet 3. In Batista de Moura and Larisse Costa for their help order for the key to include this species we propose in field. This study had the support of Fundação the following modification: de Apoio ao Desenvolvimento do Ensino, Ciência 3. Penis segment 2 with a dense patch of setae e Tecnologia do Estado de Mato Grosso do Sul on either side of midline ... 9 (FUNDECT) / Coordenação de Aperfeiçoamento

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